Sequential mate encounters: female but not male body size influences female remating behavior

Whether and how individuals choose sequentially among matesis an important but largely neglected aspect in sexual selectionstudies. Here, we explore female remating behavior in the cellarspider Pholcus phalangioides. We focus on body size as one ofthe most important traits involved in mate choice. Large andsmall females (n = 216) were double mated with large or smallmales in all eight possible combinations. All females copulatedwhen virgin, but only 82% accepted a second male. The chanceof a female remating was not significantly predicted by thebody size of the second or first male or by the size differencebetween the two. In contrast, a previous study demonstrateda male size effect in that larger males monopolized femalesuntil egg laying when two males of different sizes were present.We suggest that sequential encounters are more common undernatural conditions than male monopolization of females becauseestimates of concurrent multiple paternity together with observationsin a natural population do not favor mate guarding as the predominantmating strategy in this species. It follows from our study thatthe intensity of sexual selection on male size may be greatlyoverestimated when using a competitive laboratory setting fora species in which females generally encounter mates in a sequentialfashion. Female remating probability was significantly predictedby female size, with large females remating with higher probabilitythan small females. Thus, when mating with large females, malesmay gain higher fertilization success through increased femalefecundity but also face a higher sperm competition risk.     

Reproductive versus ecological advantages to larger body size in female snakes, Vipera aspis

Body size can influence an organism's microevolutionary fitness either via ecological factors (ecological selection) or changes in reproductive output (sexual or fecundity selection). Published studies on sexual dimorphism in reptiles have generally focussed on sexual-selective forces on males, under the implicit assumption that the intensity of fecundity selection in females (and hence, overall selection on female body size) is likely to be relatively consistent among lineages. In this paper, we explore the degree to which larger body size enhances ecological attributes (e.g., food intake, growth, survival) and reproductive output (reproductive frequency, litter size, offspring size, offspring viability) in free-ranging female aspic vipers, Vipera aspis . The less-than-annual reproductive frequency of these animals allows us to make a direct comparison between females in years during which they concentrate on "ecological" challenges (especially building energy reserves) versus reproductive challenges (producing a litter). Because female snakes have limited abdominal space to hold the clutch (litter), we expect that fecundity should depend on body size. However, our data show that larger body size had a more consistent effect on ecological attributes (such as feeding rates and "costs of reproduction") than on reproductive output per se. Indeed, total reproductive output was maximised at intermediate body sizes. These results suggest that variation in female body size among and within species (and hence, in the degree of sexual dimorphism) may be driven by the ecological as well as reproductive consequences of body size variation in both sexes.     

Sexual selection for large male size in a polyandrous butterfly: the effect of body size on male versus female reproductive success in Pieris napi

Sexual selection theory predicts that the larger sex shouldbe that for which fitness increases at the faster rate withsize. In butterflies, as in most invertebrates, females areusually the larger sex, but previous comparative analysis hasshown that relative male size increases with female polyandryamong butterflies. In agreement with this pattern, males arelarger than females in the strongly polyandrous green-veinedwhite butterfly, Pieris napi L., and in this article we assessthe size dependence of reproductive success in both sexes. Inan experiment where virgin males and females were released inthe field, we found no strong association between size and malemating success. However, laboratory experiments showed thatthere was a strong correlation between size and the ejaculatethat the male delivered to the female at mating and that largeejaculates delayed female remating for a longer time comparedto small ejaculates. Moreover, female P. napi utilize male-derivednutrients received at mating to increase their fecundity. Hence,large males sire more offspring both by way of donating morenutrients to female egg production and by way of delaying femaleremating (given that the last male to mate with the female willfather most of the offspring). Laboratory experiments showedthat the association between size and fecundity was low, ornonexistent, among P. napi females allowed to mate only once.However, weak size dependence was found for polyandrous females.We hypothesize that size dependence of female fecundity maybe especially weak among polyandrous butterflies because a fundamentalsource of variation in fecundity relates to their ability tofind nutrient giving males, an ability which may be unrelatedto female size. According to this hypothesis there is a causalassociation between weak size dependence of female fecundityand polyandry, and a strong size dependence of male reproductivesuccess that may underlie the comparative pattern of positivecorrelation between relative male size and polyandry.     

The influence of male and female body size on copulation duration and fecundity in Drosophila melanogaster

We studied two components of the mating system, copulation duration and early fecundity, in relation to body size in Drosophila melanogaster. Body size variation was created experimentally by varying the degree of crowding (starvation) among larvae from an inbred strain, keeping the genetics and temperature as constant as possible. Hence, in contrast to most previous studies, where genetic and environmental variation have been confounded, we aimed at investigating how much pure phenotypic variation could influence copulation duration and early fecundity. It is shown that copulation duration and fecundity both strongly dependent on female body size, but either not or much less so on male body size. Small females mate faster than medium or large females and small females have the lowest fecundity. Among males, medium-size males are more fecund than smaller or larger males, resulting in stabilising selection for intermediate male size. These results are in contrast with previous findings.     

Male steroid hormones and female preference for male body odor

It has been suggested that human scent works as a signal in mate selection, but the empirical evidence is scarce. Here, we examined whether women's olfactory preferences for a man's scent could be correlated with his testosterone, estradiol, or cortisol concentrations, and whether these preferences change along with the menstrual cycle. In line with previous studies, women in their most fertile period gave the highest attractiveness ratings to all men. However, the intensity ratings by women at different menstrual phases did not significantly differ statistically. Interestingly, we found that cortisol concentration in saliva correlated positively with the attractiveness but not with the intensity ratings of male T-shirt odor by all women's groups. However, neither testosterone nor estradiol was significantly associated with the ratings of attractiveness or intensity. Thus, our study suggests that there could be a novel mechanism for odor-based selection in humans.     

The relationship between female body size and egg size in pipefishes

Comparing five species of pipefish, egg size was significantly larger in species with brood pouches (Syngnathus typhle, Syngnathus acus and Syngnathus rostellatus) than in species without brood pouches (Entelurus aequoreus and Nerophis ophidion). Egg size correlated positively with female body size in species with brood pouches, but was similar across female sizes in the species lacking pouches. These results may reflect differences in offspring competition as a consequence of variable offspring relatedness within a brood, due to the mating systems adopted by the different species and the presence or absence of a brood pouch.     

Field evidence that host selection by conopid parasitoids is related to host body size

Summary The body size of the host insect in which a parasitoid develops can have important effects on its development and life history. Large and small host body size have both been suggested to be advantageous to parasitoids, depending on the life-history of the species concerned. We test field data on the bumblebeeBombus terrestris and its conopid parasitoids for evidence of differences in size between parasitised and unparasitised worker bees. Bees acting as hosts for conopid parasitoids are on average larger-bodied than unparasitised bees. This result holds for bees collected in two different years, and whether bees are collected while foraging or from the nest. The results we present demonstrate differential parasitism of hosts of different body sizes, but do not necessarily indicate active host choice by conopids. However, they are in agreement with independent evidence that conopids develop more successfully in large-than in small-bodied hosts.     

Empirical evidence for an optimal body size in snakes

Abstract The concept of optimal size has been invoked to explain patterns in body size of terrestrial mammals. However, the generality of this phenomenon has not been tested with similarly complete data from other taxonomic groups. In this study we describe three statistical patterns of body size in snakes, all of which indicate an optimal length of 1.0 m. First, a distribution of largest body lengths of 618 snake species had a single mode at 1.0 m. Second, we found a positive relationship between the size of the largest member of an island snake assemblage and island area and a negative relationship between the size of the smallest member of an island snake assemblage and island area. Best-fit lines through these data cross at a point corresponding to 1.0 m in body length, the presumed optimal size for a one-species island. Third, mainland snake species smaller than 1.0 m become larger on islands whereas those larger than 1.0 m become smaller on islands. The observation that all three analyses converge on a common body size is concordant with patterns observed in mammals and partial analyses of four other disparate animal clades. Because snakes differ so strikingly from mammals (ectotherms, gape-limited predators, elongate body shape) the concordant patterns of these two groups provide strong evidence for the evolution of an optimal body size within independent monophyletic groups. However, snakes differ from other taxonomic groups that have been studied in exhibiting a body size distribution that is not obviously skewed in either direction. We suggest that idiosyncratic features of the natural history of ectotherms allow relatively unconstrained distributions of body size whereas physiological limitations of endotherms constrain distributions of body size to a right skew.     

Determinants of male ejaculate investment in the cotton bollworm Helicoverpa armigera: mating history,female body size and male age

Many studies demonstrate that ejaculate size may be influenced by male condition, female quality and the risk or intensity of sperm competition. In the present study, the effect of male and female conditions, male mating history and female mating status on ejaculate sperm numbers in the polyandrous moth Helicoverpa armigera is examined. A large variation in ejaculate size is found and, although female body size and male age influence ejaculate size, female age and copula duration do not. Both male and female mating histories have significant effects on ejaculate sperm numbers. Males reduce ejaculate expenditure in successive matings but deliver significantly more apyrene and eupyrene sperm to nonvirgin than to virgin females.     

The effect of female body size on male mounting behaviour in Dermacentor variabilis and D. andersoni

Abstract. This study was undertaken to determine if body size of female ticks was an important factor in eliciting the mating behaviour of Dermacentor variabilis and D. andersoni males. Dummy female ticks (DFTs) representing varying sizes of feeding females were prepared from plastic beads. Size of DFT was shown to be important in determining both the time spent by males in contact with DFTs and the extent of male mating response released. Dermacentor varlabilis males preferred the smaller-sized DFT, whereas D. andersoni males preferred the larger sizes. Males of both species were able to discriminate between size of DFT only when mounting sex pheromone (MSP) was present.
Size of the DFT was more important than the composition of the MSP extract in determining the time males spent in contact with DFTs.
Males of both species were sensitive to variations in concentration of extract, and spent 2–6 times longer in contact with DFTs when the concentration was optimal than when it was not. No significant difference in the male's mating response was seen in response to variation in concentration of heterospecific extract.
Male mating response, i.e. the behaviour of the male as it progresses through the initial contact, climbing onto the dorsal surface and turning onto the venter, was shown not to be dependent upon how long the male spent in contact with the DFT. Male D. andersoni had a much lower mating response to DFTs than D. variabilis males, suggesting that some further stimulus may be required.     

Effects of female body size and age and male mating status on male combat in Anastatus disparis (Hymenoptera: Eupelmidae)

1. Aggressive behaviour is widely exhibited by animals to acquire important resources and usually shows a dangerous or nondangerous pattern. Dangerous fighting patterns are usually characterized by fights ending with contestants being severely injured or killed. Resource value is an important nonstrategic factor influencing fighting behaviour. Studies of many species addressing nondangerous fighting behaviour have shown that when resource values change, organisms usually adjust their fighting behaviour accordingly. Only a few species show dangerous fighting patterns. Thus, few relevant studies have addressed how variation in resource value affects aggression with a dangerous fighting pattern. 2. Here, an egg parasitoid wasp, Anastatus disparis, which exhibits a dangerous fighting pattern to acquire mating opportunities, was used as an experimental model to study the adjustment of fighting behaviour resulting from a change in resource value. 3. Our results show that the female properties of body size and age affect their objective resource value and that males increased their fighting intensity for relatively large and young females. However, male mating status in A. disparis may not influence the subjective value of mate resources, and fighting intensity did not significantly differ between mated and virgin males. In addition, the number of times a male had previously mated had no significant effect. These results suggest that mating opportunities are important for both virgin and mated males, resulting in neither of them showing any adjustment in fighting for mating opportunities. 4. Generally, A. disparis males with extreme fighting patterns adjust their fighting behaviour according to the variation in resource value, which avoids the meaningless costs of injury and death.     

Thermal body patterns for healthy Brazilian adults (male and female)

The aim of this study was to establish the skin temperature (T_sk) thermal profile for the Brazilian population and to compare the differences between female and male Brazilian adults. A total of 117 female and 103 male were examined with a thermographic camera. The T_sk of 24 body regions of interest (ROI) were recorded and analyzed. Male T_sk results were compared to female and 10 ROI were evaluated with respect to the opposite side of the body (right vs. left) to identify the existence of significant contralateral T_sk differences (ΔT_sk). When compared right to left, the largest contralateral ΔT_sk was 0.3 °C. The female vs. male analysis yielded significant differences (p<0.05) in 13 of the 24 ROI. Thigh regions, both ventral and dorsal, had the highest ΔT_sk by sex (≈1.0 °C). T_sk percentile below P₅ or P₁₀ and over P₉₀ or P₉₅ may be used to characterize hypothermia and hyperthermia states, respectively. Thermal patterns and T_sk tables were established for Brazilian adult men and women for each ROI. There is a low T_sk variation between sides of the body and gender differences were only significant for some ROIs.     

Nuptial gifts and the evolution of male body size

In many insect systems, males donate nuptial gifts to insure an effective copulation or as a form of paternal investment. However, if gift magnitude is both body size-limited and positively related to fitness, then the opportunity exists for the gift to promote the evolution of large male size. In the striped ground cricket, Allonemobius socius, males transfer a body size-limited, somatic nuptial gift that is comprised primarily of hemolymph. To address the implications of this gift on male size evolution, we quantified the intensity and direction of natural (fecundity) and sexual (mating success) selection over multiple generations. We found that male size was under strong positive sexual selection throughout the breeding season. This pattern of selection was similar in successive generations spanning multiple years. Male size was also under strong natural selection, with the largest males siring the most offspring. However, multivariate selection gradients indicated that gift size, and not male size, was the best predictor of female fecundity. In other words, direct fecundity selection for larger gifts placed indirect positive selection on male body size, supporting the hypothesis that nuptial gifts can influence the evolution of male body size in this system. Although female size was also under strong selection due to a size related fecundity advantage, it did not exceed selection on male size. The implications of these results with regard to the maintenance of the female-biased size dimorphic system are discussed.     

It is not always about body size: evidence of Rensch's rule in a male weapon

In many species, sexual dimorphism increases with body size when males are the larger sex but decreases when females are the larger sex, a macro-evolutionary pattern known as Rensch''s rule (RR). Although empirical studies usually focus exclusively on body size, Rensch''s original proposal included sexual differences in other traits, such as ornaments and weapons. Here, we used a clade of harvestmen to investigate whether two traits follow RR: body size and length of the fourth pair of legs (legs IV), which are used as weapons in male–male fights. We found that males were slightly smaller than females and body size did not follow RR, whereas legs IV were much longer in males and followed RR. We propose that sexual selection might be stronger on legs IV length than on body size in males, and we discuss the potential role of condition dependence in the emergence of RR.     

Empirical evidence for fast temperature-dependent body size evolution in rotifers

Organisms tend to decrease in size with increasing temperature by phenotypic plasticity (the temperature-size rule; ectotherms) and/or genetically (Bergmann’s rule; all organisms). In this study, the evolutionary response of body size to temperature was examined in the cyclically parthenogenetic rotifer Brachionus plicatilis. Our aim was to investigate whether this species, already known to decrease in size with increasing temperature by phenotypic plasticity, presents a similar pattern at the genetic level. We exposed a multiclonal mixture of B. plicatilis to experimental evolution at low and high temperature and monitored body size weekly. Within a month, we observed a smaller size at higher temperature, as compared to body size at lower temperature. The pattern was consistent for the size of both mature females and eggs; rotifers kept at high temperature evolved to be on average 14% (after 2 weeks) and 3% (after 3 weeks) smaller than the ones kept at low temperature (10 and 5% in the case of eggs, respectively). We therefore found that B. plicatilis is genetically programmed to adjust its body size-to-environmental temperature.     

Seasonal trends in body size of adult male mosquitofish,Gambusia affinis,with evidence for their social control

Synopsis Collections from a natural population of mosquitofish,Gambusia affinis, in southern Indiana showed that males born early in the breeding season mature rapidly at a small body size, while males born later in the breeding season delay maturation and achieve larger body sizes. A field experiment, involving removal of males from pond populations, was conducted to test the hypothesis that delayed maturation by late summer males is, at least in part, under social control. Mean total length (TL) of adult and maturing males in late summer in ponds from which large numbers of males had previously been removed was significantly lower than mean TL of males in control ponds. These data support the hypothesis of social control and support Sohn's competition-predation hypothesis for control of adult male body size inGambusia.     

Male contest investment changes with male body size but not female quality in the spider Nephila clavipes

Animals use rules to adjust their level of investment in a contest. We evaluate male strategies during contests over females in the golden orb-web spider Nephila clavipes. We tested whether male behaviour changes with female value, and found that contests were similar in intensity and outcome whether the female was a juvenile or adult, virgin or non-virgin, or whether one male had invested sperm in the female. We found evidence that males use a self-assessment strategy when deciding to withdraw from a contest. Loser body size and contestant size difference were correlated with a higher frequency of contest escalation, and fights involving two large males were more likely to escalate than a fight in which one male was small. A multiple regression showed that loser body size had a stronger effect on contest escalation than contestant body size difference. More importantly, the size of the winning male had no effect on contest escalation, a key prediction of a self-assessment strategy. In N. clavipes, body size is the primary factor that determines the outcome of male contests, and males do not appear to assess their opponent or the quality of the resource when deciding to withdraw from the fight.